Fossil Ants

Fossil ants include both specimens fossilised in shale or other stone (compression fossils) and ants embedded in Amber. Copal is a general term that refers to non-fossilised resin material whatever its geological age. Copal older than Holocene is referred to as fossil copal.

The geologic age of the deposits are important in understanding the evolution of ants as well as their extinction and the distribution of ants worldwide. In some cases the surrounding vegetation and insects that are preserved along with the ants allows a partial reconstruction of the habitat and in some cases the microhabitat as well.

Winged and worker ants were much larger in times past so that some ant specimens are still identifiable as ants when found in stone. Specimens preserved in amber are in much better condition allowing for a more reliable identification.

Photo Gallery

One of the earliest known ants, Gerontoformica cretacica, from Charentese amber, Aquitaine Basin, France. It belongs to the subfamily Specomyrminae, the most primitive group of true ants.
Azteca alpha from Dominican Republic Amber.
An inclusion from Baltic amber.
Protazteca elongata, based on an impression fossil.

Amber - Fossil Resins

A few plant resins have the ability to fossilize and insects trapped and embeded in this matrix are known as amber fossils. Resin that is older than one million years are designated as amber and younger aged resins are known as copal.

Amber is often designated by the type of plant exudate or given a mineral name. Ant fossils are most often named for its geographical origin; Dominican amber, Mexican amber, Baltic amber, Saxonian amber, Rovno amber, Spanish amber, Lebanese amber, etc.

Copal

Subfossil resins or Copal can be Carbon-14 dated so that the resin sold as "amber" in Madagascar by gem dealers is approximately 50 years old. The Copal from Colombia has been dated as 250 years old.

Ichnofossils

Trace fossils of ant nests. The evidence for these fossils representing preserved ant nest structures are based on interpretation of various elements of the ichnofossils that are analogous to modern day ant nests.

Fossil Taxa

Subfamilies
Genera
Species
All fossils (subfamilies, genera and species)

Geologic Time Zones Containing Ants

Pliocene - About 5.333 to 3.6 million years ago.
Miocene - About 23.03 to 5.332 million years ago.
Oligocene - About 34 million to 23 million years before the present.
Eocene - Lasted approximately from 56 to 34 million years ago.
Paleocene - Lasted from about 65.5 to 56 million years ago.
Cretaceous - From circa 145.5 to 65.5 million years ago.

Taxa by Geologic Time Zones

Era	Period	Epoch	Age	Start, million years age	Formations	Taxa
Cenozoic	Quaternary	Holocene	Meghalayan	0.0042
			Northgrippian	0.0082
			Greenlandian	0.0117
		Pleistocene	Late ('Tarantian')	0.126
			Chibanian	0.781
			Calabrian	1.8
			Gelasian	2.58
	Neogene	Pliocene	Piacenzian	3.6	No formations known	No taxa known
		Pliocene	Zanclean	5.333	Lac Chambon, France Fossil	Formica pitoni • Lasius chambonensis
		Miocene	Messinian	7.246	Brunn-Vösendorf, Austria Fossil • Varvara Formation Fossil	Camponotites ullrichi • Paraphaenogaster pannonica • Polyrhachis annosa
			Tortonian	11.63	Auxillac, France Fossil • Epecuen Formation Fossil • Joursac, France Fossil • Montagne d'Andance, France Fossil • Schossnitz, Poland Fossil • Sicilian Amber	Attaichnus kuenzelii • Camponotites crozei • Camponotites obesus • Carebara schossnicensis • Carebara sophiae • Cataulacus planiceps • Cataulacus silvestrii • Ectatomma gracile • Formica auxillacensis • Formica cantalica • Formica maculipennis • Hypopomyrmex bombiccii • Incertogaster praecursor • Lasius crispus • Lasius oblongus • Leptomyrmula maravignae • Lonchomyrmex nigritus • Oecophylla grandimandibula • Oecophylla sicula • Plagiolepis labilis • Podomyrma mayri • Ponera leptocephala • Sicelomyrmex corniger • Tapinoma minutissimum • Technomyrmex deletus
			Serravallian	13.82	Berezovsky Massif Fossil • Khronya Cape Fossil • Oeningen, Switzerland Fossil • Ormety, Georgia Fossil • Parschlug, Austria Fossil	Camponotites heracleus • Camponotites induratus • Camponotites lignitus • Carebara rugiceps • Formica demersa • Formica globiventris • Formica gravida • Formica immersa • Formica lavateri • Formica longicollis • Formica macrophthalma • Formica martynovi • Formica orbata • Formica primitiva • Formica primordialis • Formica procera • Formica pulchella • Formica seuberti • Imhoffia nigra • Imhoffia pallida • Lasius occultatus • Lasius tertiarius • Liometopum crassinervis • Liometopum escheri • Liometopum globosum • Liometopum longaevum • Liometopum stygium • Liometopum venerarium • Liometopum ventrosum • Myrmica angusticollis • Myrmica minor • Myrmica molassica • Myrmica obsoleta • Paraphaenogaster aemula • Paraphaenogaster macrocephala • Solenopsis atavina
			Langhian	15.97	Chon-Tyz Mine Fossil • Iki Island, Japan Fossil • Kuban Province Fossil • Malyi Kamyshlak Fossil • Peruvian Amber • Shanwang, China Fossil • Vishnevaya Balka Creek Fossil • Zhangpu Amber	Alloiomma changweiensis • Alloiomma differentialis • Camponotites ambon • Camponotites ampullosus • Camponotites curviansatus • Camponotites gracilis • Camponotites longus • Camponotites microthoracus • Camponotites pictus • Camponotites plenus • Camponotites shanwangensis • Camponotites xiejiaheensis • Casaleia inversa • Dolichoderus evolans • Dolichoderus jiaoyanshanensis • Dolichoderus lacinius • Dolichoderus luridivenosus • Dolichoderus tavridus • Dolichoderus transversipetiolaris • Elaphrodites mutatus • Elaphrodites scutulatus • Eurymyrmex geologicus • Formica ceps • Formica flavifemoralis • Formica linquensis • Formica ovala • Iridomyrmex shandongicus • Kotshkorkia laticeps • Lasius inflatus • Lasius mordicus • Lasius truncatus • Lasius validus • Lasius vetulus • Leptogenys lacerata • Liometopum eremicum • Liometopum lubricum • Liometopum palaeopterum • Liometopum potamophilum • Miosolenopsis fossilis • Myopopone sinensis • Odontoponera pseudotransversa • Oecophylla macroptera • Oecophylla taurica • Pachycondyla minutansata • Pachycondyla nubeculata • Paraphaenogaster avita • Paraphaenogaster dumetorum • Paraphaenogaster lapidescens • Paraphaenogaster microphthalma • Paraphaenogaster paludosa • Paraphaenogaster shanwangensis • Ponerites taurica • Ponerites umbra • Rhytidoponera kirghizorum • Taphopone karaganensis • Taphopone stauropolitana • Tapinoma baculum • Tapinoma neli • Technomyrmex septentrionalis • Zhangidris atopogaster
			Burdigalian	20.44	Barstow Formation Fossil • Borneo Amber • Chon-Tyz Mine Fossil • Iki Island, Japan Fossil • Kuban Province Fossil • Lake Victoria, Kenya Fossil • Malyi Kamyshlak Fossil • Mokrina, Czechia Fossil • Most Formation Fossil • Radoboj Fossil • Shanwang, China Fossil • Vishnevaya Balka Creek Fossil	Alloiomma changweiensis • Alloiomma differentialis • Attopsis longipennis • Camponotites ambon • Camponotites ampullosus • Camponotites curviansatus • Camponotites gracilis • Camponotites heracleus • Camponotites induratus • Camponotites lignitus • Camponotites longus • Camponotites microthoracus • Camponotites pictus • Camponotites plenus • Camponotites shanwangensis • Camponotites xiejiaheensis • Casaleia inversa • Casaleia longiventris • Cataulacus plebeius • Dolichoderus evolans • Dolichoderus heeri • Dolichoderus jiaoyanshanensis • Dolichoderus lacinius • Dolichoderus luridivenosus • Dolichoderus tavridus • Dolichoderus transversipetiolaris • Elaphrodites mutatus • Elaphrodites scutulatus • Emplastus antiquus • Emplastus dubius • Emplastus haueri • Emplastus miocenicus • Emplastus ocellus • Eurymyrmex geologicus • Formica aemula • Formica ceps • Formica flavifemoralis • Formica fragilis • Formica globiventris • Formica kollari • Formica lavateri major • Formica linquensis • Formica oblita • Formica obscura • Formica obtecta • Formica obvoluta • Formica oculata • Formica ovala • Formica parexsecta • Formica ungeri • Gesomyrmex bremii • Heeridris croatica • Imhoffia pallida • Iridomyrmex shandongicus • Kotshkorkia laticeps • Lasius anthracinus • Lasius globularis • Lasius inflatus • Lasius longaevus • Lasius longipennis • Lasius mordicus • Lasius occultatus • Lasius ophthalmicus • Lasius truncatus • Lasius validus • Lasius vetulus • Leptogenys lacerata • Liometopum brunascens • Liometopum croaticum • Liometopum eremicum • Liometopum imhoffii • Liometopum lubricum • Liometopum palaeopterum • Liometopum pallidum • Liometopum potamophilum • Lonchomyrmex freyeri • Miosolenopsis fossilis • Myopopone sinensis • Myrmecites latus • Myrmecites pusillus • Myrmica venusta • Odontomachus paleomyagra • Oecophylla leakeyi • Oecophylla macroptera • Oecophylla obesa • Oecophylla taurica • Pachycondyla minutansata • Pachycondyla nubeculata • Palaeosminthurus juliae • Paraphaenogaster avita • Paraphaenogaster dumetorum • Paraphaenogaster jurinei • Paraphaenogaster lapidescens • Paraphaenogaster microphthalma • Paraphaenogaster paludosa • Paraphaenogaster shanwangensis • Paraphaenogaster tertiaria • Ponerites atavinus • Ponerites elongatus • Ponerites gracilior • Ponerites nitidus • Ponerites oblongiceps • Ponerites taurica • Ponerites tenuis • Ponerites umbra • Rhytidoponera kirghizorum • Taphopone karaganensis • Taphopone stauropolitana • Tapinoma baculum • Technomyrmex septentrionalis • Zhangidris atopogaster
			Aquitanian	23.03	Decín, Czechia Fossil • Ethiopian Amber • Mae Sot, Thailand Fossil • Mexican Amber	Aphaenogaster praerelicta • Camponotites novotnyi • Cephalotes maya • Cephalotes olmecus • Cephalotes poinari • Cephalotes ventriosus • Cyphomyrmex maya • Desyopone hereon • Discothyrea maya • Forelius chenpauch • Melissotarsus ethiopiensis • Pheidole chaan • Pheidole pauchil • Proceratium poinari • Ravavy goldmani • Schismiscapus exstinctum • Technomyrmex svojtkai
	Paleogene	Oligocene	Chattian	28.1	Bitterfeld Amber • Dominican Amber • Enspel Formation Fossil • Foulden Maar Diatomite Fossil • Rott, Germany Fossil • Sumatran Amber • Velikaya Kema Fossil	Acanthognathus poinari • Acanthostichus hispaniolicus • Acropyga glaesaria • Agroecomyrmex duisburgi • Anochetus ambiguus • Anochetus brevidentatus • Anochetus conisquamis • Anochetus corayi • Anochetus dubius • Anochetus exstinctus • Anochetus intermedius • Anochetus lucidus • Anochetus miosumatrensis • Aphaenogaster amphioceanica • Aphaenogaster archaica • Aphaenogaster enspelensis • Aphaenogaster oligocenica • Aphaenogaster sommerfeldti • Apterostigma electropilosum • Apterostigma eowilsoni • Austroponera schneideri • Azteca alpha • Azteca eumeces • Bradoponera meieri • Bradoponera similis • Camponotus parvus • Carebara antiqua • Carebara kutscheri • Cephalotes alveolatus • Cephalotes bloosi • Cephalotes brevispineus • Cephalotes caribicus • Cephalotes dieteri • Cephalotes hispaniolicus • Cephalotes integerrimus • Cephalotes jansei • Cephalotes obscurus • Cephalotes serratus • Cephalotes squamosus • Cephalotes sucinus • Ctenobethylus goepperti • Cylindromyrmex antillanus • Cylindromyrmex electrinus • Cylindromyrmex inopinatus • Cyphomyrmex taino • Discothyrea gigas • Dolichoderus balticus • Dolichoderus brevipennis • Dolichoderus caribbaeus • Dolichoderus cornutus • Dolichoderus dibolius • Dolichoderus intermedius • Dolichoderus kutscheri • Dolichoderus mesosternalis • Dolichoderus passalomma • Dolichoderus primitivus • Dolichoderus prolaminatus • Dolichoderus sculpturatus • Dolichoderus tertiarius • Eldermyrmex exsectus • Electromyrmex wheeleri • Emplastus obliquus • Emplastus similis • Eocamponotus mengei • Eocenomyrma rugosostriata • Fallomyrma transversa • Formica bauckhorni • Formica flori • Formica gustawi • Formica kutscheri • Formica phaethusa • Gesomyrmex hoernesi • Gnamptogenys casca • Gnamptogenys europaea • Gnamptogenys levinates • Gnamptogenys pristina • Goniomma poschmanni • Gracilidris humiloides • Hypoponera atavia • Ilemomyrmex caecus • Lasius schiefferdeckeri • Lasius vulgaris • Leptomyrmex neotropicus • Liometopum rhenanum • Metalasius pumilus • Monomorium mayrianum • Monomorium pilipes • Mycetomoellerius primaevus • Myrmecorhynchus novaeseelandiae • Myrmica nungesseri • Myrmica paradoxa • Myrmica saxonica • Myrmica schaefersi • Neivamyrmex ectopus • Neoponera vejestoria • Nesomyrmex caritatis • Nesomyrmex dominicanus • Nylanderia pygmaea • Nylanderia vetula • Odontomachus pseudobauri • Odontomachus spinifer • Oecophylla brischkei • Oecophylla crassinoda • Oxyidris antillana • Pachycondyla succinea • Pachycondyla tristis • Paraneuretus longicornis • Paraneuretus tornquisti • Paraphaenogaster bizeri • Paraphaenogaster freihauti • Paraphaenogaster loosi • Paraphaenogaster schindleri • Paraphaenogaster ussuriensis • Paraphaenogaster wettlauferi • Paraphaenogaster wuttkei • Paraponera dieteri • Pheidole anticua • Pheidole primigenia • Pheidole tethepa • Plagiolepis klinsmanni • Plagiolepis kuenowi • Plagiolepis paradoxa • Plagiolepis solitaria • Plagiolepis squamifera • Platythyrea dentata • Platythyrea procera • Platythyrea pumilio • Platythyrea scalpra • Plesiomyrmex tubulatus • Ponera elegantissima • Ponera wheeleri • Poneracantha brunoi • Prenolepis henschei • Prionomyrmex longiceps • Prionomyrmex wappleri • Pristomyrmex archaios • Procerapachys annosus • Proceratium denticulatum • Proceratium dominicanum • Proceratium gibberum • Proceratium poinari • Protaneuretus mirabilis • Pseudomyrmex antiquus • Pseudomyrmex avitus • Pseudomyrmex baros • Pseudomyrmex coruscus • Pseudomyrmex macrops • Pseudomyrmex nexilis • Pseudomyrmex oryctus • Pseudomyrmex prioris • Pseudomyrmex succinus • Pseudomyrmex thecolor • Pseudomyrmex vicinus • Rhytidoponera gibsoni • Rhytidoponera waipiata • Stigmomyrmex rugulosus • Stigmomyrmex venustus • Strumigenys electrina • Strumigenys pilosula • Strumigenys poinari • Strumigenys schleeorum • Sussudio boreus • Tapinoma electrinum • Tapinoma troche • Technomyrmex caritatis • Technomyrmex hispaniolae • Technomyrmex kemaensis • Temnothorax gracilis • Temnothorax longaevus • Temnothorax praecreolus • Tetraponera europaea • Tetraponera lacrimarum • Tetraponera ocellata • Tetraponera simplex • Unnamed Enspel Formation Fossil Ants • Yantaromyrmex constrictus • Yantaromyrmex geinitzi • Yantaromyrmex intermedius • Yantaromyrmex samlandicus • Zatania electra
		Oligocene	Rupelian	33.9	Aix-en-Provence, France Fossil • Amgu River Fossil • Canyon Ferry Reservoir • Quesnel, British Columbia Fossil	Aphaenogaster longaeva • Calyptites antediluvianus • Camponotites longiventris • Camponotites penninervis • Camponotites theobaldi • Carebara thorali • Dolichoderus coquandi • Dolichoderus obliteratus • Emplastus amguensis • Formica arcana • Formica latinodosa • Formica sepulta • Formica serresi • Lasius epicentrus • Liometopum greenwalti • Manica iviei • Pachycondyla globiventris • Paraphaenogaster maculata
		Eocene	Priabonian	37.8	Bembridge Marls Fossil • Bolshaya Svetlovodnaya Fossil • Brunstatt, France Fossil • Celas, France Fossil • Florissant Fossil • Fonseca Formation Fossil • Kleinkems, Germany Fossil • Rovno Amber • Wellborn, Texas Fossil	Agastomyrma laticeps • Aphaenogaster antiqua • Aphaenogaster mersa • Asymphylomyrmex balticus • Biamomyrma lata • Biamomyrma rugosa • Biamomyrma zherikhini • Bilobomyrma ukrainica • Boltonidris mirabilis • Bradoponera meieri • Britaneuretus anglicus • Camponotites cockerelli • Camponotites compactus • Camponotites vehemens • Carebara antiqua • Carebara nitida • Carebara ucrainica • Casaleia orientalis • Cataglyphoides constrictus • Cataglyphoides dlusskyi • Colobopsis brodiei • Ctenobethylus goepperti • Damzenomyrmex ribbeckei • Dolichoderus affectus • Dolichoderus balticus • Dolichoderus bruneti • Dolichoderus explicans • Dolichoderus jonasi • Dolichoderus lucidus • Dolichoderus oviformis • Dolichoderus passalomma • Dolichoderus perkovskyi • Dolichoderus pilipes • Dolichoderus pinguis • Dolichoderus polessus • Dolichoderus robustus • Dolichoderus tertiarius • Dolichoderus vectensis • Dolichoderus vlaskini • Dolichoderus zherichini • Drymomyrmex rasnitsyni • Emplastus biamoensis • Emplastus britannicus • Emplastus elongatus • Emplastus gurnetensis • Emplastus hypolithus • Emplastus kozlovi • Emplastus macrops • Enneamerus reticulatus • Eocamponotus mengei • Eocenomyrma breviscapa • Eocenomyrma orthospina • Eocenomyrma rugosostriata • Eocenomyrma ukrainica • Eoformica expectans • Fallomyrma anodonta • Fallomyrma marginata • Fallomyrma robusta • Fallomyrma transversa • Fonsecahymen stigmata • Formica alsatica • Formica biamoensis • Formica eoptera • Formica flori • Formica gustawi • Formica palaeopolonica • Formica paleosibirica • Formica phaethusa • Formica radchenkoi • Formica ribbeckei • Formica tripartita • Gesomyrmex hoernesi • Gesomyrmex incertus • Gesomyrmex macrops • Gesomyrmex magnus • Glaphyromyrmex oligocenicus • Gnamptogenys europaea • Hypoponera atavia • Incertogaster primitiva • Iridomyrmex breviantennis • Lasius schiefferdeckeri • Lelejus venustus • Leucotaphus donisthorpei • Leucotaphus gurnetensis • Leucotaphus permancus • Liometopum incognitum • Monomorium kugleri • Monomorium mayrianum • Monomorium pilipes • Myrmecites dubius • Myrmecites major • Myrmecites sibiricus • Myrmica electrina • Myrmica intermedia • Myrmica rudis • Nylanderia pygmaea • Oecophylla atavina • Oecophylla megarche • Oecophylla praeclara • Oecophylla superba • Pachycondyla aberrans • Pachycondyla calcarea • Pachycondyla conservata • Pachycondyla dubia • Pachycondyla globiventris • Pachycondyla oligocenica • Pachycondyla succinea • Paraneuretus dubovikoffi • Paraneuretus tornquisti • Paraphaenogaster hooleyana • Paraphaenogaster maculipes • Plagiolepis klinsmanni • Plagiolepis kuenowi • Plagiolepis minutissima • Plagiolepis solitaria • Plagiolepis squamifera • Ponera lobulifera • Ponera mayri • Ponerites antropovi • Ponerites crawleyi • Ponerites hooleyi • Prenolepis henschei • Prionomyrmex longiceps • Pristomyrmex elmesi • Proceratium eocenicum • Proceratium petrosum • Proleptothorax primitivus • Protaneuretus succineus • Solenopsis alena • Solenopsis blanda • Solenopsis foersteri • Solenopsis maxima • Solenopsis moesta • Solenopsis privata • Solenopsis superba • Solenopsis valida • Solenopsites rossi • Sussudio boreus • Syntaphus wheeleri • Taphopone aberrans • Taphopone macroptera • Taphopone microptera • Taphopone petrosa • Tapinoma electrinum • Tapinoma glaesaria • Temnothorax gracilis • Temnothorax longaevus • Tetraponera europaea • Tetraponera ocellata • Tetraponera oligocenica • Tetraponera simplex • Vollenhovia kipyatkovi • Yantaromyrmex constrictus • Yantaromyrmex geinitzi • Yantaromyrmex mayrianum • Yantaromyrmex samlandicus • Ypresiomyrma orientalis
			Bartonian	41.2	Danish Amber • Kuclín, Czechia Fossil	Dolichoderus kutschlinicus
			Lutetian	47.8	Baltic Amber • Belarus Amber • Bournemouth Fossil • Elko, Nevada Fossil • Sakhalin Amber • Tallahatta Formation Fossil	Agroecomyrmex duisburgi • Aneuretellus deformis • Aphaenogaster dlusskyana • Aphaenogaster groehni • Aphaenogaster mersa • Aphaenogaster oligocenica • Aphaenogaster ribbeckei • Aphaenogaster sommerfeldti • Asymphylomyrmex balticus • Bilobomyrma baltica • Bradoponera electrina • Bradoponera meieri • Bradoponera wunderlichi • Carebara antiqua • Carebara groehni • Cataglyphoides constrictus • Cataglyphoides intermedius • Chimaeromyrma brachycephala • Ctenobethylus goepperti • Dissimulodorylus perseus • Dlusskyus groehni • Dolichoderus balticus • Dolichoderus brevicornis • Dolichoderus brevipalpis • Dolichoderus cornutus • Dolichoderus elegans • Dolichoderus granulinotus • Dolichoderus jonasi • Dolichoderus longipennis • Dolichoderus longipilosus • Dolichoderus mesosternalis • Dolichoderus nanus • Dolichoderus passalomma • Dolichoderus polessus • Dolichoderus polonicus • Dolichoderus punctatus • Dolichoderus robustus • Dolichoderus sculpturatus • Dolichoderus tertiarius • Dolichoderus vexillarius • Drymomyrmex claripennis • Drymomyrmex fuscipennis • Eldermyrmex oblongiceps • Electromyrmex klebsi • Electromyrmex wheeleri • Electroponera dubia • Enneamerus costatus • Enneamerus reticulatus • Eocamponotus mengei • Eocenomyrma elegantula • Eocenomyrma groehni • Eocenomyrma orthospina • Eocenomyrma rugosostriata • Eotapinoma compacta • Eotapinoma gracilis • Formica flori • Formica gibbosa • Formica gustawi • Formica heteroptera • Formica horrida • Formica lucida • Formica luteola • Formica macrognatha • Formica nigra • Formica palaeopolonica • Formica parvula • Formica phaethusa • Formica quadrata • Formica strangulata • Formica surinamensis • Formica trigona • Formica zherikhini • Formicium mirabile • Gesomyrmex hoernesi • Glaphyromyrmex oligocenicus • Gnamptogenys europaea • Gnamptogenys rohdendorfi • Hypoponera atavia • Lasius nemorivagus • Lasius punctulatus • Lasius schiefferdeckeri • Liometopum oligocenicum • Manica andrannae • Metalasius pumilus • Monomorium mayrianum • Monomorium pilipes • Myrmica damzeni • Myrmica dictyosa • Myrmica electrina • Myrmica eocenica • Myrmica intermedia • Myrmica longispinosa • Myrmica rudis • Nylanderia pygmaea • Oecophylla bartoniana • Oecophylla brischkei • Oecophylla crassinoda • Pachycondyla baltica • Pachycondyla gracilicornis • Pachycondyla succinea • Parameranoplus primaevus • Paraneuretus longicornis • Paraneuretus tornquisti • Pityomyrmex tornquisti • Plagiolepis klinsmanni • Plagiolepis kuenowi • Plagiolepis singularis • Plagiolepis solitaria • Plagiolepis squamifera • Plagiolepis succini • Plagiolepis wheeleri • Platythyrea primaeva • Ponera lobulifera • Ponera mayri • Prenolepis henschei • Prionomyrmex gusakovi • Prionomyrmex janzeni • Prionomyrmex longiceps • Procerapachys annosus • Procerapachys favosus • Procerapachys sulcatus • Proceratium eocenicum • Prodimorphomyrmex primigenius • Protaneuretus succineus • Protoformica proformicoides • Protomyrmica atavia • Protopone primigena • Pseudocamponotus elkoanus • Stenamma berendti • Stigmatomma electrinum • Stigmatomma groehni • Stigmomyrmex rugulosus • Stigmomyrmex venustus • Stiphromyrmex robustus • Sussudio boreus • Tapinoma electrinum • Temnothorax glaesarius • Temnothorax gracilis • Temnothorax hystriculus • Temnothorax longaevus • Temnothorax petiolatus • Temnothorax placivus • Tetramorium kulickae • Tetramorium paraarmatum • Tetraponera groehni • Tetraponera klebsi • Tetraponera lacrimarum • Tetraponera ocellata • Tetraponera simplex • Thanacomyrmex hoffeinsorum • Vollenhovia beyrichi • Vollenhovia prisca • Yantaromyrmex constrictus • Yantaromyrmex geinitzi • Yantaromyrmex mayrianum • Yantaromyrmex samlandicus • Zherichinius horribilis • Zherichinius rapax
			Ypresian	56	Arkansas Amber • Coldwater Beds Fossil • Eckfeld, Germany Fossil • Fushun Amber • Green River Formation Fossil • Horsefly River, British Columbia Fossil • Isle of Purbeck Fossil • Kishenehn Formation Shale Fossil • Klondike Formation Fossil • McAbee Fossil • Messel Fossil • Mo-Clay Fossil • Oise Amber • Puryear, Tennessee Fossil • Sweetwater, Wyoming Fossil • Ventana Formation Fossil	Archimyrmex piatnitzkyi • Archimyrmex rostratus • Archimyrmex smekali • Archimyrmex wedmannae • Avitomyrmex elongatus • Avitomyrmex mastax • Avitomyrmex systenus • Brachytarsites dongzhouheensis • Camponotites tokunagai • Camponotites vetus • Casaleia eocenica • Cephalopone grandis • Cephalopone potens • Clavipetiola asiatica • Curticorna leptogastrosa • Curtipalpulus eocenicus • Cyrtopone curiosa • Cyrtopone elongata • Cyrtopone microcephala • Cyrtopone striata • Dolichoderus dlusskyi • Dolichoderus kohlsi • Eoaenictites castanifurvus • Eocenidris crassa • Eoecophylla quilchenensis • Eoformica brevipetiola • Eoformica globularis • Eoformica latimedia • Eoformica magna • Eoformica pinguis • Eogorgites cingulatus • Eoleptocerites chinensis • Eoleptocerites fushunensis • Eomyrmex guchengziensis • Eoponerites longipetiolatus • Eurytarsites fushunensis • Formica annosa • Formicium berryi • Formicium brodiei • Furcisutura wanghuacunensis • Fushuniformica spinata • Fushunomyrmex liaoningensis • Gesomyrmex breviceps • Gesomyrmex curiosus • Gesomyrmex flavescens • Gesomyrmex gallicus • Gesomyrmex germanicus • Gesomyrmex miegi • Gesomyrmex pulcher • Huaxiaformica mallepetiolata • Incertogaster aurora • Iridomyrmex mapesi • Klondikia whiteae • Kohlsimyrma gracilis • Kohlsimyrma laticeps • Kohlsimyrma longiceps • Ktunaxia jucunda • Lasius glom • Leptogasteritus capricornutes • Liaoformica longipetiolata • Longicapitia reticulata • Longiformica dongzhouheensis • Longiformica guchengziensis • Macabeemyrma ovata • Magnogasterites curticornutus • Messelepone leptogenoides • Mianeuretus eocenicus • Myrmeciites goliath • Myrmeciites herculeanus • Myrmeciites tabanifluviensis • Myrmecites rotundiceps • Oecophylla eckfeldiana • Oecophylla kraussei • Oecophylla longiceps • Orbicapitia reticulata • Orbigastrula guchengziensis • Ovalicapito fushunensis • Ovaligastrula xilutianensis • Pachycondyla eocenica • Pachycondyla labandeirai • Pachycondyla lutzi • Pachycondyla messeliana • Pachycondyla parvula • Pachycondyla petiolosa • Pachycondyla petrosa • Platythyrea dlusskyi • Ponerites coloradensis • Ponerites eocenicus • Ponerites hypoponeroides • Ponerites kishenehne • Proiridomyrmex rotundatus • Proiridomyrmex vetulus • Propalosoma gutierrezae • Protazteca eocenica • Protopone dubia • Protopone germanica • Protopone magna • Protopone oculata • Protopone sepulta • Protopone vetula • Protrechina carpenteri • Pseudectatomma eocenica • Pseudectatomma striatula • Pseudomyrmex saxulum • Quadrulicapito longa • Quineangulicapito fushunensis • Sinoformica longicapitata • Sinomyrmex brunneus • Sinomyrmex furvis • Sinotenuicapito badis • Solenopsites abdita • Solenopsites minutus • Sphaerogasterites longipetiolatus • Titanomyrma gigantea • Titanomyrma lubei • Titanomyrma simillima • Wilsonia liaoningensis • Wilsonia megagastrosa • Wumyrmex furvis • Ypresiomyrma bartletti • Ypresiomyrma orbiculata • Ypresiomyrma rebekkae
		Paleocene	Thanetian	59.2	No formations known	No taxa known
			Selandian	61.6	Menat Formation Fossil	Tyrannomecia inopinata
			Danian	66	Paskapoo Formation Fossil	Napakimyrma paskapooensis
Mesozoic	Cretaceous	Late	Maastrichtian	72.1 ± 0.2	No formations known	No taxa known
			Campanian	83.6 ± 0.2	North Carolina Amber	Baikuris ocellantis • Brownimecia inconspicua • Sphecomyrma nexa
			Santonian	86.3 ± 0.5	Foremost Formation Amber • Ola Formation Fossil • Yantardakh Fossil	Archaeopone taylori • Armania curiosa • Armania robusta • Boltonimecia canadensis • Cananeuretus occidentalis • Canapone dentata • Chronomyrmex medicinehatensis • Cretomyrma arnoldii • Cretomyrma unicornis • Dlusskyidris zherichini • Eotapinoma macalpini • Haidoterminus cippus • Poneropterus sphecoides • Pseudarmania aberrans • Pseudarmania rasnitsyni
			Coniacian	89.8 ± 0.3	No formations known	No taxa known
			Turonian	93.9	Orapa Fossil	Afromyrma petrosa • Afropone oculata • Afropone orapa • Orapia minor • Orapia rayneri
			Cenomanian	100.5	Burmese Amber • Emanra Formation Fossil • Kzyl-Zhar Fossil • Raritan (New Jersey) Amber	Aquilomyrmex huangi • Archaeopone kzylzharica • Archaeopone taylori • Armania capitata • Armania pristina • Baikuris casei • Brownimecia clavata • Burmomyrma rossi • Camelomecia janovitzi • Camelosphecia fossor • Camelosphecia venator • Ceratomyrmex ellenbergeri • Ceratomyrmex planus • Chonidris insolita • Dhagnathos autokrator • Dilobops bidentata • Gerontoformica contega • Gerontoformica gracilis • Gerontoformica magna • Gerontoformica orientalis • Gerontoformica pilosa • Gerontoformica robusta • Gerontoformica rugosa • Gerontoformica spiralis • Gerontoformica sternorhabda • Gerontoformica subcuspis • Gerontoformica tendir • Haidomyrmex cerberus • Haidomyrmex davidbowiei • Haidomyrmex scimitarus • Haidomyrmex zigrasi • Kyromyrma neffi • Linguamyrmex brevicornis • Linguamyrmex rhinocerus • Linguamyrmex vladi • Myanmyrma gracilis • Myanmyrma maraudera • Protoceratomyrmex revelatus • Protozigrasimecia chauli • Sphecomyrma freyi • Sphecomyrma mesaki • Zigrasimecia boudinoti • Zigrasimecia caohuijiae • Zigrasimecia chuyangsui • Zigrasimecia ferox • Zigrasimecia goldingot • Zigrasimecia hoelldobleri • Zigrasimecia perrichoti • Zigrasimecia thate • Zigrasimecia tonsora • Zigrasimecia zui
		Early	Albian	~113	Baikura Fossil • Charentese Amber	Baikuris maximus • Gerontoformica cretacica • Gerontoformica occidentalis • Haidomyrmodes mammuthus
			Aptian	~125	Crato, Brazil Fossil	Cariridris bipetiolata
			Barremian	~129.4	No formations known	No taxa known
			Hauterivian	~132.9	No formations known	No taxa known
			Valanginian	~139.8	No formations known	No taxa known
			Berriasian	~145	No formations known	No taxa known

Subfamily-level Phylogeny with Fossil Occurrences

A dated subfamily-level phylogeny with first confident fossil occurrences for each lineage, last occurrences indicated for extinct lineages. Fossil deposit numbers correspond with Table 2, Figure 4, and Figure 5. Tree topology as well as crown and divergence dates from MOREAU & BELL (2013), which is so far the largest Formicidae-wide analysis with respect to taxon sampling. No crown age included for Aneuretinae, Martialis, and Paraponerinae as these are monotypic. Agroecomyrmecinae includes no crown age due to insufficient terminal sampling. Lineages with dotted lines were not placed directly through analyses, but rather added to the molecule-derived topology of Moreau & Bell, which did not include fossils. Stem ants and Brownimecia were placed according to conservative results of BARDEN & GRIMALDI (2016) – Note that while Sphecomyrminae and stem group ants are depicted on a single lineage, this group is not necessarily monophyletic. Formiciinae placement is based on WARD (2007), although this position was not recovered in the only two phylogenetic treatments of the subfamily (BARONI URBANI & al. 1992, GRIMALDI & al. 1997), it seems plausible to this author (Barden, 2017, Fig. 3).

Fossil Formations Known to Contain Ants

Formation	No. of Species	Start Age	End Age	Period	Epoch	Age	Locality	Country	Coordinates	Paleocoordinates
Aix-en-Provence, France Fossil	11	28.4	23.0	Paleogene	Oligocene	Chattian		France	43.5°N, 5.5°E
Amgu River Fossil	1	33.9	28.1	Paleogene	Oligocene	Rupelian	Amgu River, Primorye	Russia	45.8°N, 137.7°E	47.5°N, 134.7°E
Arkansas Amber	3	48.6	40.4	Paleogene	Eocene	Lutetian	Arkansas	United States	34.4°N, 92.8°W
Auxillac, France Fossil	2	7.5	5.3	Neogene	Miocene	Messinian		France	45.1°N, 2.9°E	45.3°N, 2.3°E
Baltic Amber	146	47.8	41.3	Paleogene	Eocene	Bartonian	Baltic Sea region	Europe	54.9°N, 19.9°E
Barstow Formation Fossil	1	19.3	13.4	Neogene	Miocene	Burdigalian to Langhian	California	United States	34.9° N, 116.8° W	34.1° N, 111.3° W
Belarus Amber	1	47.8	41.3	Paleogene	Eocene			Belarus	52.1°N, 24.2°E
Bembridge Marls Fossil	23	37.2	33.9	Paleogene	Eocene	Priabonian	Isle of Wight	United Kingdom	50.7° N, 1.4° W
Berezovsky Massif Fossil	1	12.7	11.6	Neogene	Miocene	Serravallian		Ukraine	48.3°N, 23.5°E
Bitterfeld Amber	67	25.3	23.8	Paleogene	Eocene	Bartonian	Baltic Sea region	Europe	51.6°N, 12.4°E
Bolshaya Svetlovodnaya Fossil	23	37.2	33.9	Paleogene	Eocene	Priabonian		Russia	46.6°N, 138.1°E
Borneo Amber	1	17.0	16.0	Neogene	Miocene			Borneo	2.7°N, 113.8°E
Bournemouth Fossil	3	47.8	41.3	Paleogene	Eocene	Bartonian	Bournemouth, Dorset	United Kingdom	50.7°N, 2.1°W
Brunn-Vösendorf, Austria Fossil	2	7.2	5.3	Neogene	Miocene		Brunn-Vösendorf	Austria	48.1°N, 16.3°E
Brunstatt, France Fossil	9	37.2	33.9	Paleogene	Oligocene	Rupelian	Brunstatt, Haut-Rhin	France	47.7°N, 7.3°E	47.1°N, 3.8°E
Burmese Amber	42	99.7	94.3	Cretaceous	Late Cretaceous	Early Cenomanian	Burmese amber, Kachin State	Myanmar	26.4°N, 96.7°E
Canyon Ferry Reservoir	2	32.1	31.9	Tertiary	Oligocene	Rupelian	Canyon Ferry Reservoir, Montana	United States	46.6°N, 112.1°W	48.2°N, 103.8°W
Celas, France Fossil	2	37.2	33.9	Paleogene	Eocene	Priabonian	Célas, Gard	France	44.1°N, 4.2°E
Charentese Amber	4	105.3	99.7	Cretaceous	Early Cretaceous	Albian	Charentese amber, Aquitaine Basin	France	45.9°N, 0.7°W	34.3°N, 2.4°E
Chon-Tyz Mine Fossil	3	16.0	11.6	Neogene	Miocene		Chon-Tyz mine, Naryn Province	Kyrgyzstan	42.2°N, 75.5°E	42.6°N, 75.0°E
Coldwater Beds Fossil	1	55.8	48.6	Paleogene	Eocene	Ypresian	Quilchena, British Columbia	Canada	50.1°N, 120.5°W	54.8°N, 103.5°W
Crato, Brazil Fossil	1	122.5	112.6	Cretaceous	Early Cretaceous	Aptian	Crato	Brazil	7.1° S, 39.7° W	8.6° S, 8.0° W
Decín, Czechia Fossil	1	23.0	16.0	Neogene	Miocene		Decín	Czechia	50.8°N, 14.2°E
Dominican Amber	86	26.0	15.0	Neogene	Miocene	Burdigalian		Dominican Republic	19.6°N, 70.8°W
Eckfeld, Germany Fossil	3	49.0	44.0	Paleogene	Eocene	Lutetian	Eckfeld	Germany	50.1°N, 6.8°E
Elko, Nevada Fossil	1	46.3	38.6	Neogene	Miocene	Priabonian	Elko, Nevada	United States	41.1°N, 115.5°W
Emanra Formation Fossil	2	94.3	89.3	Cretaceous	Late Cretaceous	Turonian	Khetana River, Khabarovsk	Russia	59.4° N, 140.3° E	66.7° N, 127.2° E
Enspel Formation Fossil	10	28.0	23.0	Paleogene	Oligocene	Chattian	Rhineland-Palatinate	Germany	50.6°N, 7.9°E
Epecuen Formation Fossil	1	9.0	6.8	Neogene	Miocene	Huayquerian	Epecuen Formation, Salinas de Hidalgo, La Pampa Prov.	Argentina	36.7°S, 64.3°W
Ethiopian Amber	4	23.0	16.0	Neogene	Miocene	Priabonian		Ethiopia
Fonseca Formation Fossil	1	37.2	23.0	Paleogene	Oligocene		Fonseca, Minas Gerais State	Brazil	20.2°S, 43.3°W
Foremost Formation Amber	6	84.9	70.6	Cretaceous	Late Cretaceous	Campanian	Alberta	Canada	49.8° N, 111.7° W
Foulden Maar Diatomite Fossil	4	23.3	19.0	Neogene	Miocene	Aquitanian		New Zealand	45.5°S, 170.2°E
Fushun Amber	36	55.8	48.6	Paleogene	Eocene	Ypresian	Liaoning	China	41.8°N, 123.9°E
Green River Formation Fossil	17	53.5	48.5	Paleogene	Eocene	Lutetian	Colorado	United States	39.5° N, 108.0° W
Horsefly River, British Columbia Fossil	1	55.0	48.0	Paleogene	Eocene	Ypresian	Horsefly River, British Columbia	Canada	52.3° N, 121.3° W	57.0° N, 103.5° W
Iki Island, Japan Fossil	1	16.0	13.7	Neogene	Miocene	Langhian	Chôjaburu, Iki Island	Japan	33.8°N, 129.8°E	35.4°N, 127.3°E
Isle of Purbeck Fossil	1	48.0	37.0	Paleogene	Eocene		Isle of Purbeck (Durlston Bay), Dorset	United Kingdom
Joursac, France Fossil	3	11.6	7.2	Neogene	Miocene		Joursac, Auvergne	France	45.1°N, 3.0°E
Khronya Cape Fossil	1	12.7	11.6	Neogene	Miocene	Sarmatian	Khronya Cape, Kerch, Crimea	Russia	45.4° N, 36.6° E
Kishenehn Formation Shale Fossil	12	48.4	38.6	Paleogene	Eocene	Lutetian	Montana	United States	48.4°N, 113.7°W
Kleinkems, Germany Fossil	16	37.3	33.9	Paleogene	Oligocene	Priabonian	Kleinkems	Germany	47.7°N, 7.5°E
Klondike Formation Fossil	3	55.8	48.6	Paleogene	Eocene	Ypresian	Republic, Washington	United States	48.7°N, 118.7°W
Kuban Province Fossil	1	16.0	11.6	Neogene	Miocene		Kuban Province, Caucasus	Russia	44.9°N, 40.6°E
Kuclín, Czechia Fossil	1	39.1	32.0	Paleogene	Eocene		Kuclín, near Bílina	Czechia	50.5°N, 13.8°E
Kzyl-Zhar Fossil	2	94.3	89.3	Cretaceous	Late Cretaceous	Turonian	Kzyl-Zhar	Kazakhstan	44.4°N, 67.3°E
Lac Chambon, France Fossil	2	5.3	3.6	Neogene	Pliocene		Lac Chambon, Auvergne	France	45.6°N, 2.9°E
Lake Victoria, Kenya Fossil	1	20.4	16.0	Neogene	Miocene		Lake Victoria	Kenya	0.5°S, 34.0°E
Malyi Kamyshlak Fossil	3	16.0	13.6	Neogene	Miocene		Malyi Kamyshlak, Kerch, Crimea	Russia	45.5° N, 36.5° E	45.7° N, 42.7° E
McAbee Fossil	8	55.8	48.6	Paleogene	Eocene	Ypresian	McAbee, British Columbia	Canada	50.8°N, 121.1°W
Menat Formation Fossil	1	61.0	60.0	Paleogene	Paleocene	Selandian	Menat, Puy-de-Dôme	France	46.104444°N, 2.905278°E
Messel Fossil	29	48.6	40.4	Paleogene	Eocene	Lutetian	Messel	Germany	49.9°N, 8.8°E
Mexican Amber	12	23.0	16.0	Neogene	Miocene		Chiapas	Mexico	16.3°N, 92.4°W
Mo-Clay Fossil	1	55.8	54.1	Paleogene	Eocene	Ypresian	Stolleklint and Manhøj, Isle of Fur	Denmark	56.9°N, 8.8°E
Montagne d'Andance, France Fossil	2	8.7	5.3	Neogene	Miocene	Turolian	Montagne d'Andance, Saint-Bauzile, Ardèche	France	44.7°N, 4.7°E
Most Formation Fossil	1	20.0	16.0	Neogene	Miocene	Burdigalian		Czechia	50.6°N, 13.7°E
North Carolina Amber	3	83.5	70.6	Cretaceous	Late Cretaceous	Early Campanian	Neuse River, North Carolina	United States	35.4°N, 78.0°W	35.6°N, 49.5°W
Oeningen, Switzerland Fossil	28	12.7	11.6	Neogene	Miocene	Messinian	Oeningen	Switzerland	47.7°N, 8.9°E
Oise Amber	3	55.0	52.0	Paleogene	Eocene	Ypresian		France	49.3°N, 2.7°E
Ola Formation Fossil	6	84.9	70.6	Cretaceous	Late Cretaceous	Early/Lower Campanian	Tenkinskii	Russia	60.2° N, 150.5° E	65.5° N, 133.8° E
Orapa Fossil	5	91.8	89.3	Cretaceous	Late Cretaceous	Turonian		Botswana
Ormety, Georgia Fossil	1	12.7	11.6	Neogene	Miocene	Serravallian	Ormety	Georgia	42.0°N, 41.9°E
Parschlug, Austria Fossil	5	13.7	12.7	Neogene	Miocene	Serravallian	Parschlug	Austria	47.5°N, 15.3°E
Paskapoo Formation Fossil	1	61.7	56.8	Paleogene	Paleocene	Tiffanian (Selandian to Thanetian)	Alberta	Canada	52.4°N, 113.8°W	58.7°N, 90.8°W
Peruvian Amber	1	15.0	12.0	Neogene	Miocene		Tamshiyacu	Peru
Puryear, Tennessee Fossil	1	48.6	40.4	Paleogene	Eocene	Priabonian	Puryear, Tennessee	United States	36.4°N, 88.3°W
Quesnel, British Columbia Fossil	4	33.9	23.0	Neogene	Miocene		Quesnel, British Columbia	Canada	53.0°N, 122.5°W
Radoboj Fossil	48	20.4	16.0	Neogene	Miocene	Burdigalian	Radoboj	Croatia	46.2°N, 15.9°E
Raritan (New Jersey) Amber	5	94.3	89.3	Cretaceous	Late Cretaceous	Turonian	New Jersey	United States	40.4°N, 74.3°W
Rott, Germany Fossil	5	23.8	24.2	Paleogene	Oligocene	Chattian	Rott, Westphalia	Germany	50.7°N, 7.3°E	50.6°N, 4.3°E
Rovno Amber	84	37.2	33.9	Paleogene	Eocene	Priabonian	Baltic Sea region	Europe	50.6°N, 26.3°E
Sakhalin Amber	8	47.8	41.3	Paleogene	Paleocene	Thanetian		Ukraine	47.4°N, 142.8°E
Schossnitz, Poland Fossil	3	11.6	5.3	Neogene	Miocene		Schossnitz (= Sosnica?), Silesia	Poland	51.0°N, 16.8°E
Shanwang, China Fossil	45	16.0	11.6	Neogene	Miocene		Shanwang	China	36.6°N, 118.7°E
Sicilian Amber	14	11.6	5.3	Neogene	Miocene	Priabonian		Italy		37.1° N, 14.8° E
Sumatran Amber	1	25.0	15.0	Neogene	Miocene		Sinamar (probably), Dharmasraya Regency, West Sumatra	Indonesia	1.4°N, 99.6°E
Sweetwater, Wyoming Fossil	1	49.5	49.5	Paleogene	Eocene	Ypresian	Sweetwater County, Wyoming	United States	42.2° N, 109.4° W	46.1° N, 95.5° W
Varvara Formation Fossil	1	7.2	5.3	Neogene	Miocene	Messinian	Apomarma, Messara Basin, Crete	Greece	35.1°N, 24.9°E
Velikaya Kema Fossil	6	28.1	23.03	Paleogene	Oligocene	Chattian	Velikaya Kema, Primorye	Russia	45.5°N, 137.2°E	46.7°N, 134.7°E
Ventana Formation Fossil	2	48.6	40.4	Paleogene	Eocene	Lutetian	Confluencia, Neuquen region	Argentina	41.1°S, 70.8°W	43.6°S, 61.1°W
Vishnevaya Balka Creek Fossil	5	16.0	11.6	Neogene	Miocene	Priabonian	Vishnevaya Balka Creek, Stavropol	Russia	45.1°N, 42.3°E
Wellborn, Texas Fossil	1	37.2	33.9	Paleogene	Eocene		Mossy Creek, near Wellborn, Texas	United States	30.5° N, 96.3° W
Willershausen, Germany Fossil	2	3.6	2.6	Neogene	Pliocene		Willershausen, Lower Saxony	Germany	51.8°N, 10.1°E
Yantardakh Fossil	3	85.0	85.0	Cretaceous	Late Cretaceous	Santonian	Yantardakh, Siberia	Russia
Zhangpu Amber	1	15.97	13.82	Neogene	Miocene	Langhian	Zhangpu County, Fujian Province	China	24.2°N, 117.9°E	24.7°N, 116.6°E

References

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